Ptychadena Boulenger, 1917

Marques, Mariana P., Ceríaco, Luis M. P., Blackburn, David C. & Bauer, Aaron M., 2018, Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola Atlas of Historical and Bibliographic Records (1840 - 2017), Proceedings of the California Academy of Sciences 65, pp. 1-501 : 129-143

Genus Ptychadena Boulenger, 1917

Ptychadena anchietae (Bocage, 1867) ANCHIETA’S GRASS FROG Rana anchietae Bocage 1867a:843 , fig. 1. Syntypes: MBL T.9.134 [3 specimens] (collector J.A. d’Anchieta),

destroyed by fire 18 March 1978. Type locality: “Benguella” [= Benguela] Benugela Province Angola. Rana anchietae : Peters (1877a:618). Rana mascareniensis : Boulenger (1882:52), Bocage (1895a:160). Rana oxyrhynchus oxyrhrynchus : Loveridge (1957:340). Ptychadena anchietae : Perret (1976a:19, 1979:18), Poynton and Broadley (1985b:146), Frost (1985:471,

2016), Poynton and Haacke (1993:14), Ruas (1996:25, 2002:144), Largen (2001:335), Channing

(2001:319), Channing et al. (2012:305), Ceríaco et al. (2016b:43).

Ptychadena (Ptychadena) anchietae : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: This broadly distributed species is known from Ethiopia south to Angola and southeast to South Africa.

Ocurrences in Angola (Map 78): The species occurs in both western regions and eastern regions near the Zambian border.

Cabinda: “Chinchoxo” [-5.10000, 12.10000]

( Peters 1877a:618). Kwanza Norte: “Dondo”

[-9.68333, 14.43333] (Poynton and Haacke

1993:14). Malanje: “Cangandala National

Park” [-9.84606, 16.72233] (Ceríaco et al.

2016b:43). Moxico: “ Diolo lake ” [-11.50000,

22.01667] ( Ruas 1996:25, 2002:144). Benguela: “Benguella” [-12.58333, 13.41667] ( Bocage 1867a:843; Loveridge 1957:340; Perret

1976a:19; Poynton and Broadley 1985b:146;

Frost 1985:471, 2016; Largen 2001:335);

“Dombe” [-12.95000, 13.10000] ( Bocage

1895a:160; Ruas 1996:25). Undetermined

Locality: “savanna from Ethiopia to Natal,

across to

Angola ” ( Cei 1977:16). MAP 78. Distribution of Ptychadena anchietae in Angola.

Taxonomic and distributional notes:

Perret (1979) erroneously referred to “ Huíla ” as the type locality. Bocage (1895a) and Boulenger (1882) considered R. anchietae to be a synonym of Rana mascareniensis (Duméril and Bibron, 1841) . The latter species was also confused with Rana oxyrhynchus (Smith, 1849) because of its morphological similarity (Perret 1976a). Poynton (1964a) recognized P. anchietae as valid, but later ( Poynton 1970) considered it to be a synonym of P. superciliaris (Günther, 1858) . Perret (1976a), however, disagreed noting that P. superciliaris is a forest species distinct from P. anchietae , and this interpretation was followed by Poynton and Broadley (1985b).

Ptychadena ansorgii ( Boulenger, 1905) ANSORGE’S GRASS FROG

Rana Ansorgii Boulenger 1905:107 , pl. 4, fig. 1. Holotype: BMNH 1947.2 .29.48 (collector W.J. Ansorge).

Type locality: “Between Benguella and Bihé” [= between Benguela and Bié], Angola. Rana ansorgii : Loveridge (1933:371, 1936a:95, 1936b:419, 1953a:373, 1957:343). Rana (Ptychadena) ansorgii : Monard (1937a:52, 1938:110), Parker (1939:142). Ptychadena ansorgei : Laurent (1950:14). Rana ansorgei : Inger (1959:541), Schmidt and Inger (1959:61). Ptychadena ansorgi : Cei (1977:16, 17). Ptychadena ansorgii : Loveridge (1936a:95), Poynton and Broadley (1985b:152), Ruas (1996:26), Largen

(2001:177), Channing (2001:321), Frost (1985:471, 2016). Ptychadena (Ptychadena) ansorgii : Frétey et al. (2011:41).

Global conservation status ( IUCN): Least Concern.

Global distribution: The species is known from Angola, southern Democractic Republic of Congo, northern Zambia, and Malawi.

Ocurrences in Angola (Map 79): The species occurs in the extreme northeast and in the western regions of the country. Lunda Norte:

“ Muita (Luembe E)” [-7.80000, 21.45000]

( Laurent 1950a:14; Ruas 1996:26). Lunda

Sul: “Dala” [-11.03333, 20.20000] ( Monard

1937a:52, 1938:110; Ruas 1996:26). Kwanza

Sul: “Congulu” [-10.86667, 14.28333] ( Parker

1936:142; Ruas 1996:26). Huambo: “Bimbi”

[-11.81667, 15.83333] ( Monard 1937a:52,

1938:110; Ruas 1996:26); “ Mt. Moco ”

[-12.41667, 15.18333] ( Parker 1936:14; Ruas

1996:26). Benguela: “ Benguela ” [-12.58333,

13.41667] ( Loveridge 1936a:95). Huíla :

“Kalukembé” [-13.78333, 14.68333] ( Monard

1937a:52, 1938:110; Ruas 1996:26). Cuando

Cubango: “ Kandingu (Kuluï)” [-14.66667,

17.70000] ( Monard 1937a:52, 1938:110; Ruas

1996:26). Undetermined Locality: “Between

Benguella and Bihé” ( Boulenger 1905:107; MAP 79. Distribution of Ptychadena ansorgii in Angola. Loveridge 1933:371, 1936b:419, 1953a:373, 1957:343; Schmidt and Inger 1959:61; Poynton and Broadley 1985b:152; Frost 1985:471, 2016; Ruas 1996:26), “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes: Relatively little is known of the biology of this species, which is found in the forests and savannas of south-central Africa.

Ptychadena bunoderma (Boulenger, 1907) ROUGH GRASS FROG

Rana bunoderma Boulenger 1907a:214 . Holotype: BMNH 1947.2 .2.86 (collector W.J. Ansorge). Type locality: “Caconda,” Huíla Province, Angola.

Rana (Ptychadena) buneli : Monard (1937a:55, 1938:114). Syntypes: MNHNC [three specimens] (collector A. Monard). Type locality: “environs de Dala,

sur le Tyiumbwé,” Lunda Sul Province, Angola.

Rana bunoderma : Schmidt (1936:129).

Ptychadena bunoderma : Laurent (1964a:142), Cei

(1977:16), Poynton and Broadley (1985b:152),

Frost (1985:471, 2016), Ruas (1996:26), Largen

(2001:177), Channing (2001:324).

Ptychadena (Ptychadena) bunoderma : Frétey et al.

(2011:41).

Global conservation status (IUCN):

Least Concern.

Global distribution: The species is known from Angola and northwestern Zambia.

Ocurrences in Angola (Map 80): The species occurs in central and eastern Angola.

Lunda Sul: “Alto Chicapa” [-10.88333,

19.23333] ( Laurent 1964a:142; Ruas 1996:26);

“Alto Chicapa, humidiherbosa des sources de

MAP 80 Distribution of Ptychadena bunoderma in la Kamassaka (Rio Camassaca)” [-10.88333, Angola.

19.25000] ( Laurent 1964a:142; Ruas 1996:26); “Alto Chicapa, humidiherbosa des sources de la Kamutongola (Rio Camuntongola)” [-10.53000, 19.15000] ( Laurent 1964a:142; Ruas 1996:26); “environs de Dala, sur le Tyiumbwé (Rio Chiumbe)” [-11.03333, 20.20000] ( Monard 1937a:55, 1938:114; Ruas 1996:26). Bié: “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:129; Ruas 1996:26). Huíla: “Caconda” [-13.73333, 15.06667] ( Boulenger 1907a:214; Poynton and Broadley 1985b:152; Frost 1985:471, 2016; Ruas 1996:26). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16).

Taxonomic and distributional notes: Gaston de Witte initially identified Monard’s types of R. buneli as “ Rana n. sp., voisine de Rana mascareniensis D.B.,” although based on Monard’s (1937a) description it is closer to P. bibronii or P. bunoderma . Laurent (1964a) provided some new records for Ptychadena bunoderma in Angola, including from near the type locality of P. buneli which he placed in the synonymy of P. bunoderma . Channing (2001) provided several records further north for the species without detailed information.

Ptychadena grandisonae Laurent, 1954 GRANDISON’S GRASS FROG

Ptychadena grandisonae Laurent 1954b:11 , pl. 1, figs. 2, 5, pl. 3, figs. 1-2, pl. 4, figs. 1, 9. Holotype: presumably MD 506 ? (collector M. de Petchkowsky). Type locality: “Muita, Luembe E” [= Luembe] Lunda

Norte, Angola. Rana (Ptychadena) bibroni : Monard (1937a:51, 1938:109). Ptychadena bibronii : Laurent (1950a:14). Ptychadena grandisonae : Laurent (1964a:139), Cei (1977:16), Poynton and Broadley (1985b:150), Frost

(1985:472, 2016), Poynton and Haacke (1993:14), Channing (2001:325). Ruas (2002:144). Rana grandisonae : Schmidt and Inger (1959:70). Ptychadena (Ptychadena) grandisonae : Frétey et al. (2011:41). Ptychadena cf. grandisonae : Conradie et al. (2016:17).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from the Democratic Republic of Congo, Angola, and Zambia.

Ocurrences in Angola (Map 61): The species occurs in the northeast of the country,

although there are some records further south.

Lunda Norte: “environs du Dundo”

[-7.36667, 20.83333] ( Laurent 1964a:139);

“ Muita (Luembe E)” [-7.80000, 21.45000]

( Laurent 1950a:14, 1954b:11; Schmidt and

Inger 1959:70; Poynton and Broadley

1985b:150; Frost 1985:472, 2016). Lunda Sul:

“Alto Chicapa, mare Tchá-Mutuka, près des sources du Cuango-Muqué” [-10.76667,

19.20000] ( Laurent 1964a:139); “Alto Chicapa, sources du Cuílo” [-10.86667, 19.40000]

( Laurent 1964a:139); “sources de la Kamutongola, Alto Chicapa” [-10.53000, 19.15000]

( Laurent 1964a:139); “Lunda” [-10.96667,

20.06667] ( Monard 1937a:51, 1938:109).

Moxico: “Luso” [-11.78333, 19.91667] (Ruas MAP 81. Distribution of Ptychadena grandisonae in Angola.

2002:144). Benguela: “Ebanga” [-12.73333,

14.73333] ( Monard 1937a:51, 1938:109). Huíla: “Kalukembé” [-13.78333, 14.68333] ( Monard 1937a:51, 1938:109); “ 5 km W of Calucembe” [-13.80000, 14.68333] (Poynton and Haacke 1993:14); “Nuntechite lagoon” [-15.13333, 13.41667] (Poynton and Haacke 1993:14). Cuando Cubango: “on the edge of the upper Longa River floodplain (53b)” [-14.56322, 18.44394] ( Conradie et al. 2016:9, 12, 17). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16).

Taxonomic and distributional notes: Laurent (1954b) considered Monard’s usage of the name Ptychadena bibroni (non Hallowell) to refer to Ptychadena grandisonae Laurent, 1954 . We agree with that conclusion and we also consider Rana (Ptychadena) bibroni ( Monard 1937 a, 1938) a synonym of grandisonae , since the distribution range for the real Ptychadena bibroni (Hallowell, 1845) is limited to the north of Gulf of Guinea, from the Gambia and Mauritania to the northeastern Democratic Republic of Congo and presumably to South Sudan ( Frost, 2016).

Ptychadena guibei Laurent, 1954 GUIBE’S GRASS FROG

Ptychadena chrysogaster Laurent 1954b:18 , pl. 2, fig. 2, pl. 1, fig. 3, pl. 3, figs. 5-6, pl. 4, figs. 3, 5. Holotype: MRAC 109096 fide Lang (1990:12). Type locality: “Lac Karago, Terr. de Kisenyi, Ruanda.” Ptychadena chrysogaster guibei Laurent 1954b:23 . Holotype: MD 2097 (collector M. de Petchkowsky). Type locality: “Muita, Luembe E” [= Luembe] Lunda Norte, Angola. Ptychadena bibroni : Laurent (1950a:14). Rana chrysogaster guibei : Schmidt and Inger (1959:65). Ptychadena chrysogaster guibei : Laurent (1964a:136), Cei (1977:16). Ptychadena guibei : Poynton and Broadley (1985b:154), Ruas (1996:26), Largen (2001:177), Channing

(2001:326), Frost (2016), Ceríaco et al. (2016b:47), Conradie et al. (2016:18). Ptychadena chrysogaster : Frost (1985:471). Ptychadena (Ptychadena) guibei : Frétey et al. (2011:41). Ptychadena (Ptychadena) chrysogaster : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from the Democratic Republic of Congo and eastern Angola, through Zambia, Botswana, Zimbabwe, Malawi, and northern Mozambique.

Ocurrences in Angola (Map 82): The species occurs especially in eastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1964a:136; Ruas 1996:26); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950:14, 1954b:23; Schmidt and Inger 1959:65; Poynton and Broadley 1985b:154; Ruas 1996:26). Moxico: “Cazombo” [-11.88333, 22.91667] ( Laurent 1964a:136; Ruas 1996:26). Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceríaco et al. 2016b:47). Cuando Cubango: “Cuito basin (32)” [-17.04880, 19.53333] ( Conradie et al. 2016:9-10, 18); “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9-10, 18); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9-10, 18). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” ( Cei 1977:16).

MAP 82. Distribution of Ptychadena guibei in Angola.

Taxonomic and distributional notes: Poynton and Broadley (1985b) elevated P. guibei to a full species. Frétey et al. (2011:41) cited Ptychadena (Ptychadena) chrysogaster Laurent, 1954 for Angola though we believe that this should be referred to P. guibei ( Frost 2016) . New material cited by Conradie et al. (2016) extends the species distribution into extreme southeastern Angola.

Ptychadena keilingi Monard, 1937 KEILING’S GRASS FROG

Rana (Ptychadena) keilingi Monard 1937a:53 , figs. 14–16. Syntypes: MNHC 90.0829 and 90.0830 (collector A. Monard). Type locality: “Dala,” Lunda Sul, Angola.

Rana (Ptychadena) keilingi : Monard (1938:112), Marx (1959:442), Laurent (1964a:141).

Ptychadena keilingi : Forcart (1946:126), Cei (1977:16), Poynton and Broadley (1985b:154), Frost (1985:472, 2016), Ruas (1996:26), Largen (2001:177), Channing (2001:327).

Ptychadena (Ptychadena) keilingi : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from the Democratic Republic of Congo and adjacent northeastern Angola to extreme northwestern Zambia.

Ocurrences in Angola (Map 83): The species occurs in northeastern Angola. Lunda Norte: “Rivière Luíta river, Poste de Cuílo (Rio

Luíta)” [-8.03333, 19.41667] ( Laurent

1964a:141; Ruas 1996:27); “Poste de Luangue,

humidiherbosa du risseau Katcheleka, affl.

ouest Luangue, entre la Lunguena et le TcháPemba (Posto do Luangue)” [-9.08333,

19.71667] ( Laurent 1964a:141; Ruas 1996:27).

Lunda Sul: “Alto Chicapa” [-10.88333,

19.23333] ( Laurent 1964a:141; Ruas 1996:27);

“Dala” [-11.03333, 20.20000] ( Monard

1937a:53, 1938:112; Forcart 1946:126; Marx

1959:442; Poynton and Broadley 1895b:154;

Frost 1985:472, 2016; Ruas 1996:26). Moxico:

“rives du lac Calundo (Lago Calundo)”

[-11.80000, 20.86667] ( Laurent 1964a:141;

Ruas 1996:27). Undetermined Locality:

“areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16). MAP 83. Distribution of Ptychadena keilingi in Angola.

Taxonomic and distributional notes: Two of the five type specimens noted by Monard (1937a) remain in the Musée d’Histoire Naturelle , La-Chaux-de Fond, Switzerland. Little is known of this frog species .

Ptychadena cf. mascareniensis (Duméril and Bibron, 1841) MASCARENE GRASS FROG

Rana mascareniensis Duméril and Bibron 1841:350 . Syntypes: MNHN 4379–4381 fide Guibé (1950), MCZ A-1044 (exchange from MNHN) fide Barbour and Loveridge (1929), USNM 10975 (2 specimens) fide Cochran (1961) (collectors J.R.C. Quoy and J.P. Gaimard, J.-J. Dussumier, J.- B. Leschenault, Nivoy ). Type locality: “îles Mascareignes, ou les Séchelles, Maurice et Bourbon.” Restricted to “Island of Réunion, Mascarene Islands (Indian Ocean)” by Cochran (1961) and “Ile Bourbon” [= Réunion] by Blommers-Schlösser and Blanc (1991).

Ptychadena mascareniensis hylaea Schmidt and Inger 1959:83 ). Holotype: FMNH 57965 (collector H.A. Beatty). Type locality: “Mount Nimba, Liberia ” Liberia.

Rana mascareniensis : Boulenger (1905:107).

Rana (Ptychadena) mascareniensis : Monard (1937a:50, 1938:108).

Rana mascareniensis mascareniensi s: Loveridge (1957:342), Ruas (1996:25, 2002:144).

Ptychadena mascareniensis : Cei (1977:16, 17), Frost (1985:473), Channing (2001 329), Pickersgill (2007a:128), Frost (2016), Conradie et al. (2016:18) .

Ptychadena mascareniensis bibroni : Perret (1979:6).

Ptychadena hylaea : Lamotte (1967:647), Pickersgill (2007a:134).

Ptychadena (Ptychadena) mascareniensis : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: Populations from throughout sub-Saharan Africa currently referred to this species likely represent multiple undescribed species or should be referred to other described species in this genus.

Ocurrences in Angola (Map 84): The species is widespread across the country.

Lunda Sul: “Dala” [-11.03333, 20.20000]

( Monard 1937a:50, 1938:108, Ruas 1996:25).

Moxico: “ Rio Caluando ” [-11.46667,

17.70000] ( Ruas 1996:25, 2002:144); “ Lago

Dilolo” [-11.50000, 22.01667] ( Ruas 1996:25,

2002:144); “ Lago Cameia ” [-11.71667,

20.80000] ( Ruas 1996:25, 2002:144). Bié:

“Chitau” [-11.43333, 17.15000] (Schmidt and

Inger 1959:84; Ruas 1996:25). Huíla: “ Huíla ”

[-11.71667, 20.80000] (Schmidt and Inger

1959:84). Cunene: “Chimporo” [-16.03333,

17.15000] ( Monard 1937a:50, 1938:108; Ruas

1996:25). Cuando Cubango: “Cuabango basin (29)” [-17.87291, 19.83333] ( Conradie et al. 2016:9 -10,18); “Cuando basin (39)”

[ -17.46333, 22.86638] ( Conradie et al. 2016:9 - MAP 84. Distribution of Ptychadena cf. mascareniensis in Angola.

10,18); “Cubango basin (41a)” [-17.46777,

23.06667] ( Conradie et al. 2016:9 -10,18); “Cubango basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9 -10,18); “Cubango basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9 -10,18); “Cuito basin (63)” [-17.93611, 21.10269] ( Conradie et al. 2016:9,12,18). Undetermined Locality: “Between Benguella and Bihé (Entre Benguela e Bié)” ( Boulenger 1905:107; Ruas 1996:25); “ Angola Ouest?” ( Perret 1979:9); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “arid territories along the coast” ( Cei 1977:17); “plateaus regions” ( Cei 1977:17).

Taxonomic and distributional notes: Ptychadena mascareniensis is a complex of multiple cryptic forms, possibly representing as many as ten separate species ( Vences et al. 2004; Zimkus et al. 2017) with the nominotypical form likely being restricted to Madagascar, the Seychelles, and the Mascarenes. While Zimkus et al. (2017) did not include Angolan populations in their molecular phylogenetic study, it is likely that Angolan populations correspond to both P. nilotica and one undescribed species (their OTU 6) that occurs in Gabon and Democratic Republic of Congo. Schmidt and Inger (1959) described Ptychadena mascareniensis hylaea from Mount Nimba, Liberia and remarked on an individual (CNHM 74214) from “Huíla in the highlands of southern Angola,” deposited in the Chicago Natural History Museum (now the Field Museum of Natural

History), that they considered to belong to hylaea . Lamotte (1967) subsequently considered hylaea as a distinct species, whereas Perret (1979) considered it to be a synonym of P. macareniensis bibroni (Hallowell, 1845) (= Ptychadena bibroni ) ( Frost 1985). Frétey et al. (2011) synonymized hylaea with P. mascareniensis , while authors such as Pickersgill (2007a) continue to recognize the species. A taxonomic revision to determine the status of this species in the country is being conducted by R. Ernst ( Conradie et al. 2016).

Ptychadena cf. mossambica ( Peters, 1854) MOZAMBIQUE GRASS FROG

Rana mossambica Peters 1854:626 . Syntypes: ZMB 4418 , 4419 a-b, 37555 (collector W.C.H. Peters). Type locality: “Cabeceira, Queillimane, Tette, Boror” [= Cabeceira, Quelimane ( Companhia do Boror )] Mozambique.

Ptychadena cf. mossambica : Conradie et al. (2016:18) .

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Kenya south to Mozambique and

KwaZulu-Natal, eastwards to Botswana, the

Caprivi Strip of Namibia and adjacent southeastern Angola.

Ocurrences in Angola (Map 85): Possibly limited to the southeastern region of the country. Cuando Cubango: “lower Cuando

River near the village of Jamba (38)”

[-17.58830, 22.65694] ( Conradie et al. 2016:9 -

10,18); “lower Cuando River near the village of Jamba (39)” [-17.46333, 22.86638] ( Conradie et al. 2016:9 -10,18); “ lower Cuando

River near the village of Jamba (43)”

[-17.53500, 23.18916] ( Conradie et al. 2016:9 -

10,18).

Taxonomic and distributional notes:

Conradie et al. (2016) recently collected some MAP 85. Distribution of Ptychadena cf. mossambica in Angola.

individuals from Cuando Cubango Province and provisionally assigned them to the Ptychadena mossambica complex.

Ptychadena oxyrhynchus (Smith, 1849) SHARP-NOSED GRASS FROG

Rana oxyrhynchus Smith 1849a : pl. 77, figs. 2, 2a, 2b, 2c and two accompanying unumbered pages. Lectotype: BMNH 58-11-25-97 designated by Guibé and Lamotte (1961:382) (collector A. Smith). Type locality: “ Cape of Good Hope” fide Guibé and Lamotte (1961”1960”:382), South Africa. According to Boulenger (1882:52) the specimens noted are from “ Cape of Good Hope” and “Natal”, as also reported by Guibé and Lamotte (1961 “1960”:382). The original type locality is “Kaffir land and the country about Port Natal.”

Rana oxyrhyncha : Bocage (1866a:53, 1870:68, 1895a:159, 1897b:210), Ferreira (1900a:53, 1904:112, 1906:160), Schmidt (1936:129), Schmidt and Inger (1959:91).

Rana oxyrhynchus : Boulenger (1882:51, 1905:108), Bocage (1887c:211),

Rana (Ptychadena) oxyrhynchus : Monard (1937a:49, 1938:107), Parker (1936:142).

Rana oxyrhynchus oxyrhynchus : Mertens (1937a:19, 1938a:427), Laurent (1950a:14, 1954a:73).

Rana (Ptychadena) oxyrhynchus oxyrhynchus : Hellmich (1957a:27).

Ptychadena oxyrhynchus : Guibé and Lamotte (1960 “1961”:382), Laurent (1964a:133), Cei (1977:17), Frost (1985:474, 2016), Ruas (1996:25), Channing (2001:334), Ceríaco et al. (2016b:46).

Ptychadena (Ptychadena) oxyrhynchus : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is widely distributed in savannas across western, Central, and into southern Africa.

Ocurrences in Angola (Map 86): The species occurs from western Angola to the northeastern regions. Lunda Norte: “Dundo”

[-7.36667, 20.83333] ( Laurent 1950a:14,

1954a:73, 1964a:133; Ruas 1996:25); “Muita

(Luembe E)” [-7.80000, 21.45000] ( Laurent

1950a:14, 1954a:73; Ruas 1996:25). Lunda

Sul: “Dala” [-11.03333, 20.20000] ( Monard

1937a:49, 1938:107; Ruas 1996:25). Moxico:

“ Teixeira de Sousa ” [-10.70000, 22.23333]

( Mertens 1937a:19). Malanje: “ Duque de Bragança ” [-9.10000, 15.95000] ( Bocage

1866a:53, 1895a:159; Boulenger 1882:51,

1905:108; Ruas 1996:25); “Pungo-Andongo”

[-9.66667, 15.58333] ( Bocage 1895a:159;

Ruas 1996:25), “Cangandala National Park”

[-9.84606, 16.72233] (Ceríaco et al. 2016b:46).

Kwanza Norte: “Rio Luinha” [-9.26667, MAP 86. Distribution of Ptychadena oxyrhynchus in 14.53333] ( Ferreira 1906:160; Ruas 1996:25); Angola.

“N’dalla Tando (Dala Tando)” [-9.30000, 14.91667] ( Ferreira 1904:112; Ruas 1996:25); “Cazengo” [-9.33333, 14.76667] ( Ferreira 1904:112; Ruas 1996:25); “Mucoso” [-9.53333, 14.65000] ( Hellmich 1957a:27). Kwanza Sul: “Congulu (Congulo)” [-10.86667, 14.28333] ( Parker 1936:142; Ruas 1996:25): “Chingo” [-11.20000, 13.85000] ( Ferreira 1904:112; Ruas 1996:25); “Novo Redondo, grotte de Furna, fleuve N’Gunza (Rio N’Guanza)” [-11.20000, 13.93333] ( Laurent 1954a:73; Ruas 1996:25). Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt 1936:129; Ruas 1996:25); “Chitau” [-11.43333, 17.15000] ( Schmidt 1936:129; Ruas 1996:25). Huambo: “Bimbi” [-11.81667, 15.83333] ( Monard 1938:107; Ruas 1996:25); “ Mt. Moco ” [-12.41667, 15.18333] ( Parker 1936:142; Ruas 1996:25). Benguela: “Cahata” [-12.35000, 14.81667] ( Bocage 1895a:159; Ruas 1996:25); “Quissange” [-12.43333, 14.05000] ( Bocage 1895a:159; Ruas 1996:25); “Quindumbo” [-12.46667, 14.93333] ( Bocage 1895a:159; Ruas 1996:25); “Benguella” [-12.58333, 13.41667] ( Bocage 1895a:159; Ruas 1996:25); “Entre Rios” [-13.01667, 14.63333] ( Hellmich 1957a:27); “Cubal” [-13.03333, 14.25000] ( Mertens 1938a:427; Ruas 1996:25); “Hanha” [-13.30000, 14.20000] ( Bocage 1897b:210; Ruas 1996:25). Huíla: “Caconda” [-13.73333, 15.06667] ( Bocage 1895a:159; Ruas 1996:25); “Kalukembé” [-13.78333, 14.68333] ( Monard 1937a:49, 1938:107; Ruas 1996:25); “Sangevé” [-13.88333, 15.83333] ( Monard 1937a:49, 1938:107; Ruas 1996:25); “Indungu” [-14.81667, 16.26667] ( Monard 1937a:49, 1938:107; Ruas 1996:25); “Osi” [-15.08333, 15.41667] ( Monard 1937a:49, 1938:107; Ruas 1996:25); “Kakulakaze (Kuluї)” [-15.41667, 15.73333] ( Monard 1937a:49, 1938:107; Ruas 1996:25). Cunene: “Riusseau Mbalé (Bale)” [-15.16667, 16.75000] ( Monard 1937a:49, 1938:107; Ruas 1996:25). Cuando Cubango: “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8 -9,18); “Cuando basin (39)” [-17.46333, 22.86638] ( Conradie et al. 2016:9 -10,18); “Cuando basin (41a)”

[-17.46777, 23.06667] ( Conradie et al. 2016:9 -10,18); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9 -10,18). Undetermined Locality: without precise locality ( Bocage 1870:68; Boulenger 1900:53; Laurent 1954:73); “Rio Quando” ( Bocage 1895a:160; Ruas 1996:25); “Rio Quanza” ( Boulenger 1905:108); “Carangigo” ( Boulenger 1882:51); “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “plateaus regions” ( Cei 1977:17); “arid territories along the coast” ( Cei 1977:17).

Taxonomic and distributional notes: This species occurs throughout much of sub-Saharan Africa in areas of moist savanna, near forest edges, in secondary vegetation with tall herbaceous vegetation, and in marshy and agricultura areas (Schmidt and Inger 1959; Ruas 1996; Rödel et al. 2004; Frost 2016). It appears to be relatively common in Angola, except in the arid and semiarid areas in southern regions of the country ( Ruas 1996). According to Channing (2001) this species often occurs alongside Amietia angolensis (Bocage, 1866) .

Ptychadena perplicata Laurent, 1964 MANY-RIDGED GRASS FROG Ptychadena perplicata Laurent 1964a:136 , fig. 36. Holotype: MD 5513 (collectors A. Barros Machado and E. Luna de Carvalho ). Type locality: “Alto Chicapa, humidiherbosa des souces du Cuílo” [= Cuílo River ] Lunda Sul Province, Angola.

Ptychadena ansorgei : Laurent (1954a:74).

Ptychadena perplicata : Cei (1977:16), Frost (1985:474, 2016), Ruas (1996:27), Largen (2000:177), Channing (2001:336).

Ptychadena (Ptychadena) perplicata : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Angola and Zambia.

Ocurrences in Angola (Map 87): Published records are limited to the far north of the country. Lunda Norte: “Dundo” [-7.36667,

20.83333] ( Laurent 1954a:74, 1964a:136; Ruas

1996:27). Lunda Sul: “Alto Chicapa, humidiherbosa des souces du Cuílo (Rio Cuílo,

nascente)” [-10.88333, 19.23333] ( Laurent

1964a:136; Frost 1985:474, 2016; Ruas

1996:27, Largen 2001:177).

Taxonomic and distributional notes:

According to Ruas (1996) and Largen (2000),

P. perplicata is only known from forest gal-

leries near the type locality. However , Channing (2001) provided several records without detailed information, presumably based on museum specimens, that suggest this species is MAP 87. Distribution of Ptychadena perplicata in Angola .

more widespread in the country.

Ptychadena porosissima ( Steindachner, 1867) STRIPED GRASS FROG

Rana porosissima Steindachner 1867:18 , pl. 1, figs. 9-13. Holotype: NHMW 14772 (collector F.A.P. Bayão) fide Häupl and Tiedemann (1978:28). Type locality: “ Angola ”.

Ptychadena loveridgei Laurent 1954b:14 . Holotype: MRAC 109038 . Type locality: “ Tare , Busanza, région d’Astrida, env. 1. 800 m., Ruanda.”

Rana porosissima : Bocage (1887a:191, 1897b:211), Schmidt and Inger (1959:96), Häupl and Tiedemann

(1978:28), Häupl et al. (1994:32). Rana mascareniensis : Bocage (1866a:53), Boulenger (1882:52). Rana mascareniensis var. porosissima : Bocage (1895a:160), Ferreira (1897b:240). Rana subpunctata : Loveridge (1953a:329). Rana (Ptychadena) mascareniensis : Guibé and Lamotte (1957:978). Rana loveridgei : Loveridge (1957:343). Ptychadena porosissima : Poynton and Broadley (1985b:149), Frost (1985:474, 2016), Ruas (1996:25), Largen

(2001:342), Channing (2001:337), Channing et al. (2012:311). Ptychadena (Ptychadena) porosissima : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is widely distributed in southern and eastern Africa. Ocurrences in Angola (Map 88): The species occurs from central-west regions to northeastern Angola. Zaire: “S. Salvador do

Congo ” [-6.26667, 14.23333] ( Bocage

1887a:191, Bocage 1895a:160; Ruas 1996:25).

Kwanza Norte: “Ambaca” [-9.26667,

15.18333] ( Bocage 1895a:160; Ruas 1996:25).

Malanje: “ Duque de Bragança ” [-9.10000,

15.95000] ( Bocage 1866a:53, 1895a:160; Boulenger 1882:52; Ruas 1996:25). Lunda Norte:

“Sombo” [-8.68333, 20.95000] ( Laurent

1954b:14). Benguela: “Quibula” [-12.28333,

14.68333] ( Bocage 1895a:160; Ruas 1996:25);

“Hanha” [-13.30000, 14.20000] ( Bocage

1897b:211; Ruas 1996:25). Huíla: “ Rio Cuce

(Rio Cusse)” [-13.51667, 15.20000] ( Ferreira

1897b:240; Ruas 1996:25); “Caconda”

[-13.73333, 15.06667] ( Bocage 1895a:160; MAP 88. Distribution of Ptychadena porosissima in Ferreira 1897b:240; Ruas 1996:25). Angola.

Taxonomic and distributional notes: Ptychadena porosissima ( Steindachner, 1867) was considered a variety of Rana mascareniensis (Duméril and Bibron, 1841) by Bocage (1895a) and Ferreira (1897b) and later by Guibé and Lamotte (1957). Ptychadena loveridgei Laurent, 1954 was synonymized with P. porosissima by Schmidt and Inger (1959). Loveridge (1953a) mistakenly considered P. porosissima a synonym of P. subpunctata , a position refuted by Schmidt and Inger (1959) who considered it a distinct species. Because of the similar general appearance of many species of Ptychadena and the use of characters of limited diagnostic utility in some descriptions, many names have been misapplied to more than one species. In this case, some records allocated to the P. mascareniensis complex may belong to P. porosissima .

Ptychadena subpunctata (Bocage, 1866) SPOTTED GRASS FROG

Rana subpunctata Bocage 1866b:73 . Holotype: MBL (collector F.A.P. Bayão) not located by Perret

(1967a:21), possibly lost or destroyed by fire 18 March 1978. Type locality: “ Duque de Bragança ”

[= Calandula] Malanje Province, Angola. Rana subpunctata : Bocage (1866a:54, 1895a:161, 1897a:203), Inger (1959:541). Rana mascareniensis subpunctata : Mertens (1938a:427). Rana subpunctata : Schmit and Inger (1959:102). Ptychadena subpunctata : Laurent (1964a:134), Perret (1976a:21), Cei (1977:16, 17), Poynton and Broadley (1985b:143), Frost (1985:476, 2016), Ruas (1996:25, 2002:143), Channing (2001:340), Pickersgill (2007a:136), Conradie et al. (2016:18).

Ptychadena (Ptychadena) subpunctata : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is known from Angola, extending east into the southern Democratic Republic of Congo and Zambia, and south to northern Botswana and northern Namibia.

Ocurrences in Angola (Map 89): The species occurs from central-west to eastern areas of Angola. Malanje: “ Duque de Bragança ” [-9.10000, 15.95000] ( Bocage

1866a:54, 1866a:73, 1895a:161, 1897a:203;

Perret 1976a:21; Schmidt and Inger 1959:102;

Poynton and Broadley 1985b:143; Frost

1985:476, 2016; Ruas 1996:25). Moxico:

“ Lago Dilolo ” [-11.50000, 22.01667] ( Ruas

1996:25, 2002:143); “ Lago Cameia”

[-11.71667, 20.80000] ( Ruas 1996:25,

2002:143); “Rives du lac Calundo” [-11.80000,

20.86667] ( Laurent 1964a:134; Ruas 1996:25);

“ Chites de la Lusiavo, affl. du Zambèze ,

Calunda” [-11.86667, 23.58333] ( Laurent

1964a:134; Ruas 1996:25); “Cazombo”

[-11.88333, 22.91667] ( Laurent 1964a:134;

Ruas 1996:25). Benguela: “Cubal” MAP 89. Distribution of Ptychadena subpunctata in Angola.

[-13.03333, 14.25000] ( Mertens 1938a:427;

Ruas 1996:25). Cuando Cubango: “Cubango basin(6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8 -9,18); “Cubango basin (27)” [-15.17127, 19.19433] ( Conradie et al. 2016:8 -9,18); “Cuito basin (30a)” [-17.50875, 20.06594] ( Conradie et al. 2016:9 -10,18); “Cuito basin (30f)” [-17.52638, 20.05825] ( Conradie et al. 2016:9 -10,18); “Cuito basin (33a)” [-16.90980, 19.30769] ( Conradie et al. 2016:9 -10,18); “Cuito basin (35)” [-16.62322, 19.05352] ( Conradie et al. 2016:9 -10,18); “Cuando basin (41a)” [-17.46777, 23.06667] ( Conradie et al. 2016:9 -10,18); “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9 -10,18); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9 -10,18); “Cuito basin (56)” [-15.06275, 19.14322] ( Conradie et al. 2016:9,12,18); “Cuito basin (63)” [-17.93611, 21.10269] ( Conradie et al. 2016:9,12,18). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16); “plateaus regions” ( Cei 1977:16); “ Angola ” (Poynton and Broadley 1985b:143); “north Angola ” (Schmidt and Inger 1959:140).

Taxonomic and distributional notes: Rana subpunctata was used by Bocage (1866a) as a nomen nudum, but in the same issue of the journal, Bocage (1866b) published a valid description of the new species. According to Perret (1976a), the holotype cited by Bocage from “Duque de Bragança ” was not present in the Museu Bocage when he examined that collection.

Ptychadena taenioscelis Laurent, 1954 DWARF GRASS FROG

Ptychadena taenioscelis Laurent 1954b:25 , pl. 5, fig. 1, pl. 4, fig. 6. Holotype: MRAC 13122 (collector G.F. de Witte). Type locality: “Lukula, près de Kiambi, Tanganyika,” Democratic Republic of Congo.

Ptychadena taenioscelis : Laurent (1964a:140), Cei (1977:16), Poynton and Broadley (1985b:153), Poynton and Haacke (1993:14), Ruas (1996:26), Channing (2001:341), Frost (2016), Conradie et al. (2016:18) .

Ptychadena pumilio : Largen (2001:342).

Ptychadena (Ptychadena) pumilio : Frétey et al. (2011:41).

Global conservation status (IUCN): Least Concern.

Global distribution: The species is broadly distributed across southern, Central, and eastern Africa.

Ocurrences in Angola (Map 90): The species occurs in the east of the country.

Lunda Norte: “ Rivière Luíta, Poste de Cuílo

(Rio Luíta)” [-8.03333, 19.41667] ( Laurent

1964a:140; Ruas 1996:26). Lunda Sul: “Luachimo, 120 km au sud de Vila Henrique de Carvalho (Luachimo)” [-10.68333, 20.26667]

( Laurent 1964a:140; Ruas 1996:26); “Alto

Chicapa, partie supérieure des chutes du Cuango-Muqué (Rio Cuango-Muqué )” [-10.76667,

19.20000] ( Laurent 1964a:140; Ruas 1996:26);

“Alto Chicapa” [-10.88333, 19.23333] ( Laurent 1964a:140; Ruas 1996:26). Malanje: “20

km NW of Quimbango (Quimbango)”

[-10.76667, 17.43333] (Poynton and Haacke

1993:14; Ruas 1996:26). Moxico: “Réserve de chasse de Cameia, fosse inondé près de la route” [-11.83333, 21.00000] (Laurent MAP 90. Distribution of Ptychadena taenioscelis in Angola.

1964a:140; Ruas 1996:26). Cuando Cubango:

“Cubango basin (3)” [-14.94277, 17.71863] ( Conradie et al. 2016:8 -9,18); “Cubango basin (6b)” [-14.67458, 17.73544] ( Conradie et al. 2016:8 -9,18); “Cubango basin (19)” [-14.70213, 17.37772] ( Conradie et al. 2016:8 -9,18); “Cuito basin (30d)” [-17.51327, 20.06111] ( Conradie et al. 2016: 9 -10,18); “Cuito basin (35)” [-16.62322, 19.05352] ( Conradie et al. 2016:19 -10,18); “Cuando basin (43)” [-17.53500, 23.18916] ( Conradie et al. 2016:9 -10,18); “Cuando basin (43b)” ( Conradie et al. 2016:9, 18); “Cuando basin (44a)” [-17.57333, 23.26000] ( Conradie et al. 2016:9 -10,18); “Cuito basin (62)” [-17.50875, 20.06608] ( Conradie et al. 2016:9,12,18). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16).

Taxonomic and distributional notes: Ptychadena taenioscelis Laurent, 1954 has occasionally been treated as a synonym of Ptychadena pumilio ( Boulenger 1920) . Perret (1979) suggested that P. taenioscelis is probably a subspecies of P. pumilio and referred records of P. taenioscelis from western and Central Africa to this species. Largen (2001) noted the occurrence of P. pumilio in Angola, as did Frétey et al. (2011) who included P. taenioscelis and P. smithi Guibé, 1960 as synonyms of P. pumilio . Further studies are needed to understand whether these are two distinct species or in fact are one and the same. However, we choose to treat this as a species distinct from Ptychadena pumilio , following Poynton and Broadley (1985b), Channing (2001), Pickersgill (2007a), and Frost (2016).

Ptychadena upembae (Schmidt and Inger, 1959) UPEMBA GRASS FROG

Rana upembae Schmidt and Inger 1959:111 , fig. 50. Holotype: MRAC 1228 (collector Mission G.F. de Witte), formerly Inst. Parcs Natl. Congo Belge fide Lang (1990:13). Type locality: “Kaswabilenga, Parc National de l’Upemba, Province Katanga, Belgian Congo ” (Schmidt and Inger 1959:111), Democratic Republic of Congo.

Ptychadena upembae machadoi Laurent 1964a:134 . Holotype: MD 5364 (unnamed local collector). Type locality: “Alto Chicapa, Lunda,” Angola.

Ptychadena upembae machadoi : Cei (1977:16).

Ptychadena upembae : Poynton and Broadley (1985b:150), Frost (1985:477, 2016), Ruas (1996:26, 2002:144), Channing (2001:342).

Ptychadena (Ptychadena) upembae : Frétey et al. (2011:42).

Global conservation status (IUCN):

Least Concern.

Global distribution: The species extends across south-central African from Angola, the southern Democratic Republic of Congo and

Zambia to Mozambique.

Ocurrences in Angola (Map 91): The species occurs in eastern Angola. Lunda Sul:

“Alto Chicapa” [-10.88333, 19.23333] ( Laurent 1964a:134; Ruas 1996:25). Moxico:

“ Lago Cameia ” [-11.71667, 20.80000] ( Ruas

1996:25, 2002:144). Undetermined Locality:

“areas of forest and savanna in the north and northeast of Angola ” ( Cei 1977:16). Channing

(2001) additionnally plotted unspecific localities in a 2” map.

Taxonomic and distributional notes:

Currently P. upembae machadoi is recognized MAP 91. Distribution of Ptychadena upembae in Angola. as a synonym of Ptychadena upembae ( Channing 2001; Frétey et al. 2011, Frost 2016).

Ptychadena uzungwensis (Loveridge, 1932) UDZUNGWA GRASS FROG

Rana mascareniensis uzungwensis Loveridge 1932a:384 . Holotype: MCZ A-16626 (collector A. Loveridge).

Type locality: “Dabaga, Uzungwe Mountains, Tanganyika Territory ” [ Tanzania]. Rana mascareniensis subpunctata : Schmidt (1936:129). Ptychadena uzungwensis : Laurent (1954b:10, 1964a:139), Cei (1977:16), Poynton and Broadley (1985b:151),

Poynton and Haacke (1993:14), Frost (1985:477, 2016); Ruas (1996:26, 2002:145), Channing (2001:344),

Conradie et al. (2016:18). Rana mascareniensis uzungwensis : Loveridge (1953a:372, 1957:342). Rana uzungwensis : Schmidt and Inger (1959:117). Ptychadena subpunctata : Ruas (1996:25). Ptychadena (Ptychadena) uzungwensis : Frétey et al. (2011:42).

Global conservation status ( IUCN): Least Concern.

Global distribution: The species is known from central eastern Angola and the southeastern Democratic Republic of Congo, Zambia and Tanzania south to northern Zimbabwe.

Ocurrences in Angola (Map 92): The species occurs chiefly in eastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] ( Laurent 1954b:10). Lunda Sul: “Alto Chicapa, hemidiherbosa près de la cascade du Cuango-Muqué”

[-10.76667, 19.20000] ( Laurent 1964a:139);

“ Alto Chicapa, Lunda hemidiherbosa des sources du Cuílo” [-10.86667, 19.40000] ( Laurent

1964a:139); “Alto Chicapa” [-10.88333,

19.23333] ( Laurent 1964a:139); “Dala”

[-11.03333, 20.20000] ( Laurent 1964a:139).

Malanje: “ 20 km NW of Quimbango ”

[-10.76667, 17.43333] (Poynton and Haacke

1993:14). Moxico: “Calombe” [-11.83333,

19.93333] ( Ruas 2002:145); “Réserve de chasse de Cameia dans la steppe de Ñarikumbi”

[-11.83333, 21.00000] ( Laurent 1964a:139).

Bié: “Gauca” [-11.18333, 17.45000] ( Schmidt

1936:129; Laurent 1954b:9; Schmidt and Inger

1959:118); “Chitau” [-11.43333, 17.15000]

( Schmidt 1936:129; Laurent 1954b:9; Schmidt and Inger 1959:118). Cuando Cubango: MAP 92. Distribution of Ptychadena uzungwensis in Angola.

“ Cuito basin (27)” [-15.17127, 19.19433]

( Conradie et al. 2016:8 -9,18). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola ”( Cei 1977:16).

Taxonomic and distributional notes: Loveridge (1953a) believed that Rana mascareniensis uzungwensis Loveridge, 1932 was erroneously synonymized with Rana subpunctata (Bocage, 1866) by Schmidt (1936) and cited specimens from “Gauca” and “Chitau” in Angola as belonging to P. uzungwensis . This species is considered valid by the majority of modern authors (e.g., Poynton and Broadley 1985b; Channing 2001; Frost 2016) and according to Poynton and Broadley (1985b) it inhabits grasslands and dambos. Poynton and Haacke (1993) reported one individual from “ 20 km NW of Quimbango” collected by G. G. A. Voigt in a water-logged grassland. Channing (2001) provided several records in the western regions of the country based on previously unpublished museum records.